Diversity, Rarity and the Evolution and Conservation of the Canary Islands Endemic Flora
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Anales del Jardín Botánico de Madrid Vol. 65(1): 25-45 enero-junio 2008 ISSN: 0211-1322 Diversity, rarity and the evolution and conservation of the Canary Islands endemic flora by J. Alfredo Reyes-Betancort1, Arnoldo Santos Guerra1, I. Rosana Guma1, Christopher J. Humphries2 & Mark A. Carine2,3 1 Unidad de Botánica Aplicada, Instituto Canario de Investigaciones Agrarias, Jardín de Aclimatación de La Orotava, c/ Retama n.º 2, 38400 Puerto de La Cruz, Santa Cruz de Tenerife, Spain 2 Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom 3Author for correspondence: [email protected] Abstract Resumen Reyes-Betancort, J.A., Santos Guerra, A., Guma, I.R., Humphries, Reyes-Betancort, J.A., Santos Guerra, A., Guma, R., Humphries, C.J. & Carine, M.A. 2008. Diversity, rarity and the evolution and C.J. & Carine, M.A. 2008. Diversidad, rareza, evolución y con- conservation of the Canary Islands endemic flora. Anales Jard. servación de la flora endémica de las Islas Canarias. Anales Jard. Bot. Madrid 65(1): 25-45. Bot. Madrid 65(1): 25-45 (en inglés). The endemic vascular flora of the Canary Islands comprises over La flora vascular endémica de las Islas Canarias comprende unos 680, taxa collectively accounting for more than 50% of the total 680 táxones, lo que viene a representar más del 50% de la flora na- native flora. To investigate geographical patterns of diversity tiva. Con objeto de investigar patrones geográficos de diversidad within the endemic flora, distribution data from published en la flora endémica, se recopilaron los datos publicados que, jun- sources together with other field observation and herbarium to con otras observaciones de campo y datos de herbario, sirvieron data were used to compile a data matrix comprising the distrib- para completar una matriz de datos que abarca la distribución de utions of ca. 90% of endemic taxa scored on a 10 × 10km UTM cerca del 90% de los táxones endémicos usando cuadrículas UTM grid. WORLDMAP was then used to investigate patterns of en- de10 × 10 km. A continuación, se utilizó el programa WORLDMAP demic diversity, range size rarity (a measure of endemicity), phy- para investigar los patrones de diversidad de los endemismos, el logenetic diversity and threatened taxon richness. Endemic tax- rango del grado de rareza (una medida de endemicidad), la diver- on richness was found to be highly heterogeneous across the sidad filogenética y la riqueza en táxones amenazados. Se observó archipelago, with cells containing between one and 139 taxa que la riqueza en endemismos es muy heterogénea a lo largo del each (0.05-22.82% of endemic diversity). Patterns of variation archipiélago, con unos valores por cuadrícula que oscilan entre 1 y in range size rarity and phylogenetic diversity were found to be 139 táxones (0,05-22,82% de la diversidad de táxones endémi- largely congruent with endemic diversity, although some cells cos). Los patrones de variación del rango del grado de la rareza y la exhibited markedly higher range size rarity scores than would be diversidad filogenética resultaron ser en gran parte congruentes predicted by their endemic diversity scores. In contrast, the pat- con la diversidad en endemismos, aunque algunas cuadrículas tern of endangered taxon richness across the archipelago dif- mostraron valores mucho más altos de rareza de los que podían ser fered markedly from endemic taxon richness. Many cells in Lan- predichos dada su diversidad de endemismos. En contraste, los pa- zarote, Fuerteventura and Gran Canaria exhibit higher endan- trones de riqueza en especies amenazadas en el archipiélago difi- gered taxon richness scores than would be predicted from their rieron marcadamente de la riqueza en táxones endémicos. Muchas endemic richness scores whereas in Tenerife, El Hierro, La Palma cuadrículas de Lanzarote, Fuerteventura y Gran Canaria mostraron and La Gomera, the converse is generally true. The implications valores más altos de riqueza en especies amenazadas que las que of the results both for understanding the evolution of Canary Is- pudieran ser predichas sobre la base de su riqueza en táxones en- land endemic diversity and for the conservation of the region’s démicos, mientras que en Tenerife, El Hierro y La Gomera la regla unique and vulnerable flora are considered. fue generalmente lo contrario. Se consideran las implicaciones que estos resultados suponen para la comprensión de la evolución de la diversidad de endemismos canaria y para la conservación de su sin- gular y vulnerable flora. Keywords: Canary Islands, endemism, species richness, range Palabras clave: Islas Canarias, endemismo, riqueza en especies, size rarity, threatened species richness, phylogenetic diversity, rango del grado de rareza, riqueza en especies amenazadas, di- evolution, conservation. versidad filogenética, evolución, conservación. 26 J.A. Reyes-Betancort & al. Introduction Bramwell, 2001). Emerson & Kolm (2005) demon- strated a close correlation between the number of sin- The Canary Islands archipelago comprises seven gle island endemics and the total number of species main islands together with a number of smaller islets per island and proposed that ‘diversity begets diversi- located off the northwest coast of Africa (Fig. 1). Ac- ty’ i.e. that higher species richness on islands is a dri- cording to the most recent biogeographic classifica- ver for higher rates of diversification. Other authors tion scheme (Rivas-Martínez, 2007), the Canary Is- have challenged this explanation for the observed pat- lands are recognised as a biogeographic province tern, suggesting that species diversity and endemicity within the Canario-Madeirense Subregion (Mediter- co-vary because abiotic factors influence both in a ranean Region), with two sub-provinces distin- similar manner (e.g. Pereira & al., 2007). Whittaker & guished: the Eastern Canaries (Lanzarote and Fuer- al. (2007) proposed an alternative model to explain teventura, and the Salvage Islands of Portugal) and the heterogeneity of the flora in which both species the Western Canaries (Gran Canaria, Tenerife, La richness and speciation rate reach a maximum when Gomera, La Palma and El Hierro). an island reaches maximum topographic complexity. The endemic flora of the Canary Islands is ex- To date, however, such analyses have focussed on tremely rich, with over 680 endemic taxa currently between-island comparisons and have not taken into recognised (species and subspecies), collectively ac- account the considerable within-island heteroge- counting for more than 50% of the total native flora neity in endemic species richness. Knowledge of such (Santos-Guerra, 2001). The Canary Islands are con- intra-island patterns of diversity could conceivably sidered as a hotspot of plant diversity within the Me- shed further light onto the ecological-evolutionary diterranean global diversity hotspot (Quézel & Mé- mechanisms promoting diversification of the region’s dail, 1995) and the high levels of endemicity observed flora. in both the Canary Islands flora and fauna led Sund- From a conservation perspective, the Canary Is- seth (2005) to describe the archipelago as one of the lands endemic flora is highly vulnerable to environ- top biodiversity hotspots in the world. mental change, especially the disruptive and de- The distribution of endemic diversity within the structive alteration brought about by human inter- Canary Islands is heterogeneous. Many endemics are ference and invasive species (Bramwell, 1990). Two extremely restricted in their distribution and a num- hundred and eleven endemic spermatophytes repre- ber of areas have been identified that exhibit a con- senting more that 30% of the endemic flora, are cur- centration of highly localised endemics (Bramwell & rently included on the Canary Islands red list (Go- Fig. 1. The Canary Islands and the approximate location of some of the major geographical features in the archipelago discussed in the text. Anales del Jardín Botánico de Madrid 65(1): 25-45, enero-junio 2008. ISSN: 0211-1322 Diversity of the Canarian flora 27 bierno de Canarias, 2000) of which 168 (23% of the species richness included the considerable intro- endemic flora) are included in the Atlas y Libro Rojo duced element in the flora that is unlikely to be of in- de la Flora Amenazada de España (Bañares & al., terest for conservation purposes. In a global analysis 2004) that covers all of Spain. In order to conserve of bird distributions, Orme & al. (2005) demonstrat- the unique and threatened flora, vegetation, land- ed that there is only very limited congruence between scapes and culture of the Canary Islands, an exten- the priority areas for conservation identified when to- sive network of protected areas has been developed. tal species richness, threatened species richness and A total of 146 protected areas have been designated, endemic richness are used. Indeed, only 2.5% of covering approximately 40% of the archipelago’s to- hotspot areas are common to all three methods. For- tal area (Gobierno de Canarias, 2001; see http://www. est & al. (2007) recently investigated patterns of tax- gobcan.es/cmayot/espaciosnaturales/categorias/ on richness and phylogenetic diversity in the Cape ase.html). Biodiversity hotspot and similarly found these two However despite the importance and vulnerability measures to be uncorrelated. Phylogenetic diversity of the Canarian flora and the extent of the protected (PD) may be defined as the total length of the evolu- area network, explicit analyses of geographical pat- tionary tree that connects the taxa within a given area terns of diversity within the flora have been extreme- (Vane-Wright & al., 1991; Faith, 1992) and may be ly limited. Gaisberg & Stierstorfer (2005) investigat- considered a measure of ‘feature diversity’. Forest & ed patterns of taxon richness and diversity within El al. (2007) demonstrated that in the Cape Flora of Hierro and demonstrated that whilst endemic diver- South Africa, taxon richness is greatest in the West- sity is highest on geologically old surfaces, diversity of ern Cape but the flora of this region has a lower PD the total flora (including the introduced flora) gener- score than would be predicted by its taxon richness.